Microstructure of cheese: Processing,
technological and microbiological
considerations
Claudia I. Pereira, Ana M.P. Gomes and F. Xavier Malcata
CBQF/Escola Superior de Biotecnologia, Universidade
Catolica Portuguesa, Rua Dr. Antonio Bernardino de
(Tel.: D351 22 558 0004; fax: D351 22 509 0351;
Cheese is a classical dairy product, which is strongly judged by
its appearance and texture; hence, a renewed interest in its mi-
crostructure has been on the rise, as sophisticated techniques
of analysis become more and more informative and widely
available. Processing parameters that affect microstructure
play a dominant role upon the features exhibited by the final
product as perceived by the consumer; rational relationships
between microstructure (which includes biochemical and
microbiological indicators), and quality and safety of the
products are accordingly required. Subsequent to that extra
fundamental knowledge, technological innovations may even-
tually improve current cheesemaking processes, and permit
mechanistic design of novel ones. This review thus focuses
on recent advances pertaining to the microstructure of cheese,
and discusses them in a logical and critical manner.
Introduction
Cheese is a highly regarded food in most human cultures,
and has accordingly been present throughout ages in man-
kind daily life. The variety of cheeses currently available is
large, because a myriad of topical advances e encompassing
manufacture and ripening, have cumulatively taken place
over time. In addition to the somewhat intrinsic (and
unpredictable) variability within each cheese type, tailor-
made cheese matrices have been proposed based on specific
microstructures e which have emerged side by side with
introduction of alternative (or improved) methodologies in
cheesemaking.
Microstructure is not a static concept; it evolves instead
along the food processing chain, and eventually leads to ma-
jor transformations relative to the original microstructure of
the milk feedstock itself. This realization thus encompasses
specific molecular compositions and spatial arrangements.
* Corresponding author.
0924-2244/$ - see front matter 2009 Elsevier Ltd. All rights reserved.
doi:10.1016/j.tifs.2009.02.006
Almeida, P-4200-072 Porto, Portugal
e-mail: fxmalcata@esb.ucp.pt)
On the other hand, it should be emphasized that, besides
the supporting proteinaceous/fatty matrix, microorganisms
are also an integral part of cheese. Microbial activity pro-
duces indeed major transformations of the cheese matrix,
which will affect the final microstructure as well. Hence, it
is also crucial to elucidate microorganismematrix interac-
tions, in attempts to understand the whole picture.
Consumer acceptance of a cheese product depends
directly on its appearance, flavour and texture e which
are in turn originated by a thorough combination of micro-
biological, biochemical and technological parameters, that
affect microstructure directly or indirectly. Note that the
ultimate success of any food product relies on consumers
reactions: in fact, human perception of organoleptic charac-
teristics is closer to the consumer status at the moment of
decision than data generated by any type of analytical
instrumentation (Adhikari, Heymann, & Huff, 2003) e
despite its constraints in repeatability and objectiveness.
Texture is intrinsically related to the arrangement of
various chemical components within distinct micro- and
macrostructure levels e e.g. proteic network or fat fraction;
it is the external manifestation of such structures that even-
tually determines the uniqueness and distinctive character
of a cheese product. However, cheeses are particularly
complex systems, so full and meaningful assessment of
the effects of microstructure (and texture) upon flavour
and appearance is still incipient.
This paper discusses a number of fundamental aspects
pertaining to microstructure of cheese e and specifically
focuses on issues associated with microstructural effects
arising from technological and processing approaches.
Manipulation of physical properties of milk
A huge variety of dairy products exists at present,
depending on the deliberate alterations of the original
characteristics of milk e which always start with some
form of bulk concentration (Malcata, 1999). A set of down-
stream operations are normally applied to milk e which are
intended to alter, strengthen or create structure; they are
aimed at minimization (or even elimination) of potential
risk factors, e.g. product decay via metabolism of contam-
inating microflora. Attempts to improve yield, as well as to
design novel products with better organoleptic features,
also lead to changes at the microstructure level.
The most important unit operations in dairy food physi-
cal manipulation, that impinge upon microstructure, are
listed in Table 1, and are discussed below in further detail.
Thermal treatments
Thermal treatment is the most common operation used with
milk; this is due to the tight safety that is enforced in food
plants, including industrial cheese manufacture. Several dis-
tinct heat treatments have been successfully applied in
industrial practice e ranging from mild to severe ones. As
expected, the more severe the heat treatment, the more exten-
sive the damages brought about thereby e as the thermal
history of milk affects the properties of the coagulum, and
thus the development afterwards of microstructure; heat-
treated (and homogenized) milk may in fact lead to off-
flavours and weak clotting features (Singh & Wauguna, 2001).
Another consequence of milk heating is whey protein
denaturation, followed by aggregation; or interaction with
caseins, in the first place. The fraction of whey proteins
taken up may reach 50e70%, depending on their degree
of denaturation. A high pre-heating temperature improves
yield and stability of the final product, as well as texture
e which will become smoother if short-time heating is em-
ployed, because of increased water binding to the denatured
whey proteins (Hinrichs, 2001).
Pressure treatments
High pressure (HP) has been proposed as a suitable tech-
nology for milk treatment to substitute, or in addition to
thermal treatment. Its relatively recent development has
been associated mainly with the possibility of inactivating
undesired microorganisms and/or enzymes in milk; how-
ever, one cannot disregard the effect of such a practice
upon milk constituents themselves, and consequently on
the final characteristics of ripened cheese e beyond im-
provement of microbial safety and extension of shelf-life.
So far, the most important concern in terms of HP-induced
changes pertains to the physicochemical properties of casein
micelles and whey proteins e as HP affects intramolecular
bonds, either reinforcing or weakening them. Lopez-
Fandino, Carrascosa, and Olano (1996) reported that pressur-
ization of cheesemaking milk at 300e400 MPa for 30 min
caused an increase of 14e20% in curd weight, and a decrease
of 7.5e15% in protein loss in whey. Huppertz, Fox, de Kruif,
and Kelly (2006), Lopez-Fandino (2006) and Considine,
Patel, Anema, Singh, and Creamer (2007) have comprehen-
sively reviewed the molecular changes induced by HP in
milk proteins, whereas Trujillo, Capellas, Saldo, Gervilla,
and Guamis (2002) highlighted the principles of HP and their
implications on the final properties of dairy products.
However, a thorough description of the aforementioned
chemical and molecular mechanisms, and of the underlying
theoretical issues is not within the scope of this review e but
rather the implications of such a type of process on the struc-
tural features of cheese.
It is widely accepted that HP treatments alter milk coag-
ulation characteristics e either by reducing or increasing
the gelation time (Lopez-Fandino, 2006), or by inducing
proteolytic and lipolytic activities that promote acceleration
of cheese ripening (Guerzoni et al., 1999). Exposure to HP

Table 1. Processing applications in cheesemaking milk, and effects thereof upon microstructure of cheese

Treatment parameters Microstructural effects References

High temperatures No effect on structure of casein micelles Kim and Jimenez-Flores (1995)
Weakened clotting properties Grappin and Beuvier (1997)
Denaturation of serum proteins Heinrichs (2001)
Interactions between k-casein and whey proteins Schreiber (2001)
Interactions between milk proteins and fat membrane components Singh and Wauguna (2001)
High pressures Effect on milk protein intramolecular bonds Guerzoni et al. (1999)
Change of coagulation characteristics OReilly et al. (1999)
No effect on activity of native milk enzymes Needs et al. (2000)
Reduction of syneresis Wendin et al. (2000)
Increase of cheese yield Capellas et al. (2001)
Closer-packed structure Kheadr et al. (2002)
Acceleration of cheese ripening Huppertz et al. (2006)
Lopez-Fandino (2006)
Enzymes (TGase) Modification of rheological and renneting properties Cozzolino et al. (2003)
Prevention of coalescence between milk fat globules Hinz, Huppertz, Kulozik
Incorporation of whey proteins and Kelly (2007)
Influence on primary and secondary stages of coagulation Bonisch et al. (2008)
Membranes Reduction of rennet coagulation time Erdem (2000)
Increase in firmness of coagulum Mistry (2004)
More compact structure of casein Benfeldt (2006)
Softer texture and altered flavour characteristics, in hard- and semi-hard cheeses
increases cell membrane permeability (Malone, Shellhammer,
& Courtney, 2002), thus favouring release of intracellular
enzymes and their access to substrates. This technique
can also be used to attenuate the viability of starter bacteria
throughout ripening, and thus enhance the release of meta-
bolic compounds upon lysis (Upadhyay, Huppertz, Kelly, &
McSweeney, 2007). Furthermore, HP aids in release of Ca,
P and as1- and as2-caseins e depending on the operating
conditions, hence leading to destabilization of micelles
(Huppertz, Fox, & Kelly, 2004; Regnault, Dumay, &
Cheftel, 2006); this renders caseins more susceptible to
eventual action by proteolytic enzymes.
From a microscopic point of view, HP treatments of milk
produce a closer-packed structure in cheese, due to the re-
duction in average size of casein micelles (Needs, Stenning,
Gill, Ferragut, & Rich, 2000); the finer structure strengthens
the cheese matrix, via establishment of caseinecasein and/or
caseinefat complexes (Kheadr, Vachon, Paquin, & Fliss,
2002), thus producing firmer curds (Lopez-Fandino et al.,
1996). These findings are well documented via microscopy,
both electronic (Capellas, Mor-Mur, Sendra, & Guamis,
2001; Kheadr et al., 2002) and optical one (Capellas et al.,
2001; Wendin, Langton, Caous, & Hall, 2000).
Such a technology has been as well applied to cheese after
manufacture (Garde, Arques, Gaya, Medina, & Nunez, 2007;
Juan, Ferragut, Guamis, & Trujillo, 2008). Several authors
(OReilly et al., 2003; OReilly, Murphy, et al., 2002;
OReilly, OConnor, Murphy, Kelly, & Beresford, 2000;
OReilly, OConnor, Murphy, Kelly, & Beresford, 2002)
have studied in depth the effects of a number of cheese HP
treatments on key ripening characteristics of both Cheddar
and Mozzarella cheeses. In general, high pressures (ca.
200e400 MPa) for relatively short times (ca. 20 min)
brought about changes in the protein structure itself e which
in turn improved the functional features of the final cheese
matrix, viz. an increase in the flowability and fluidity, and a re-
duction in the melt time upon heating. On the other hand,
lower pressures (ca. 50e200 MPa) combined with longer
processing times (up to 82 h) did not seem to affect structural
features, as indicated by confocal laser scanning microscopy.
Membrane treatments
Membrane technology is more and more widely used in
the dairy industry, because of its versatility and high ener-
getic efficiency. It basically consists on a pressure-driven
operation through a porous solid medium, aimed at separa-
tion and/or concentration; it ranges from plain filtration to
reverse osmosis e also known as hyperfiltration (Mistry,
2004), depending on the percolating pore diameter and con-
sequent particle size partition capacity, from ca. 100 mm
down to ca. 0.1 nm (Kelly, 2004).
The main goal of ultrafiltration (UF) is to concentrate
bulk milk via retention of whey proteins; this enhances yield
of cheese, which also contains lower concentrations of lac-
tose and minerals (Hinrichs, 2001). A few physicochemical
properties of milk are critical in cheesemaking via UF e viz.
viscosity, buffering capacity and rennet-driven coagulation.
Upon performance of UF on milk, proteins and salts are
simultaneously concentrated, thus leading to an increased
buffering capacity; this influences metabolism of lactic
acid bacteria favourably, whereas coagulation time is re-
duced and firmness of the resulting coagulum is increased
(Mistry, 2004).
Furthermore, membrane treatments have been used for
recovery of milk components and their incorporation as
new ingredients in cheese manufacture formulations e
viz. UF retentates (Govindasamy-Lucey, Jaeggi, Bostley,
Johnson, & Lucey, 2004), milk protein concentrates and
phosphocasein (Guinee, OKennedy, & Kelly, 2006).
Therefore, membrane technology can be regarded as an
efficient tool for development and continuous production
of cheese, via intermediate production of highly concen-
trated matrices e that can be coagulated with little to no
acid whey production at all (Nelson & Barbano, 2005).
Despite its intrinsic advantages, UF has not yet been fully
adopted for manufacture of hard- and semi-hard cheeses e
because of the harder texture in the final product, when
compared with conventional soft and semi-soft cheeses.
Furthermore, the high water binding capacity of whey pro-
teins e coupled with retarded proteolysis, influences the tex-
ture of cheese, which becomes progressively firmer (i.e.
more resistant to fracturing), more cohesive, grainier and
drier, whereas the structure of the protein matrix becomes
coarser and more compact, thus resulting in slower softening
during ripening (Fox, McSweeney, Cogan, & Guinee, 2000).
Finally, Erdem (2000) suggested that casein micelles de-
crease in size, and possibly undergo rearrangement via estab-
lishment of hydrophobic bonds e thus producing a more
compact structure and an increase in elasticity; Karami, Eh-
sani, Mousavi, Rezaei, and Safari (2009) provided evidence
for this fact via scanning electron microscopy. However, fun-
damental knowledge on how the changes in milk proteins
and fat during UF affect cheese manufacturing is still limited.
Enzymatic treatments
Milk coagulation can be influenced by various factors, in-
cluding enzyme-mediated cross-linking of casein micelles
via microbial transglutaminase (TGase) e a transferase
that forms bonds both within and between several proteins,
via glutamine and lysine residues (Kashiwagi et al., 2002).
Caseins are particularly suitable substrates for this activity,
owing to their low level of tertiary structure (Huppertz &
de Kruif, 2007).
Cross-linking of food proteins brought about by TGase
affects their gelling, rheological, emulsifying and renneting
properties; Bonisch, Heidebach, and Kulozik (2008) showed
that such cross-linking affects both the primary and the
secondary stages of rennet coagulation. Hinz et al. (2007)
reported, in turn, that TGase-induced cross-linking of milk
proteins affects considerably their emulsifying features,
via increasing the stability of fat globules against coales-
cence. Finally, Cozzolino et al. (2003) suggested use of
TGase as a means to incorporate whey proteins into cheese-
making milk, in a way somewhat similar to UF. These fea-
tures seem promising in cheese manufacture, as they
increase yield and gel strength e thus favouring
microstructure.
Characterization of cheese
The whole cheesemaking process contributes to develop
a distinct and more complex matrix than the departing feed-
stock milk. During coagulation e which, in most cases, is
enzyme-driven, rennet enzymes bring about breakdown of
k-casein, that is present especially on the surface of casein
micelles. After such a chemical step, physical agglomera-
tion takes place e which gives rise to a more uniform pro-
tein mass, that spontaneously expresses whey; this process
of compactation (termed syneresis) may be taken one step
further, by externally applying pressure onto the curd. On
the other hand, fat globules normally retain their mem-
branes e and are thus observed as single entities, which
may form clusters that are entrapped within the so formed
proteinaceous, three-dimensional matrix.
Physical matrix
Cream cheese is a spreadable soft cheese e which
structurally differs from other cheeses because of its
lack of a compact protein matrix, coupled with a relatively
highmoisture content. Its major structural component is fat
(ca. 33%), in the form of clusters of globules interspersed
within milk proteins (Kalab, 1993); its microstructure has
been defined as corpuscular, or composed of compact fat-
casein aggregates with large spaces filled with whey
(Kalab, 1985). The moisture content thereof is a factor that
positively contributes to spreadability: if protein particles
and fat globules are packed less densely, a high-moisture
product will result.
An increasing demand for reduced fat cheese has arisen
among more educated, health-aware consumers. However,
decreasing the level of (or even totally removing) fat often
leads to textural drawbacks, viz. a hard body with poor
meltability and stretchability (Merrill, Oberg, McManus,
Kalab, & McMahon, 1996), as well as a rubbery texture
(Mistry, 2001). The flavour, colour and mouthfeel of cheese
will also be adversely affected, since a few compounds
dissolve preferentially in fat (Li, Marshall, Heymann, &
Fernando, 1997) e although a higher smoothness may
also be imparted to the cheese matrix (Mistry, 2001).
Fat substitutes e based on proteins, polysaccharides and
synthetic chemical molecules (or even alternative fats),
have been tested in attempts to convey at least as satisfac-
tory sensory attributes; among others, Wendin et al. (2000)
proved that it is possible to create a cream cheese with
a lower fat content, but possessing an organoleptic profile
similar to that of normal cream cheeses e simply by crite-
rious modification of processing parameters. A more recent
study (Sahan, Yasar, Hayaloglu, Karaca, & Kaya, 2008)
showed that low-fat cheeses containing fat replacers exhibit
a higher level of total free fatty acids than low-fat control
cheeses, with a significant increase in their texture attri-
butes and meltability; however, the full-fat cheese was
awarded the best sensory scores, at all stages of ripening.
Scanning electron micrographs helped elucidate how total
(or partial) substitution of regular milk fat produces cheeses
with different microstructures and inherently distinct tex-
tural characteristics (Lobato-Calleros et al., 2007).
Another good example of a matrix for which the interre-
lationship between microstructure and final quality is
apparent is processed cheese. Cheese processing was orig-
inally proposed as a means to upgrade cheese of lower
quality. Processed cheeses are typically prepared by heating
a mixture of comminuted cheeses in the presence of salts
(Piska & Stetina, 2004) e which usually include sodium
citrate or phosphate, the main role of which is sequestering
calcium from the calciumecaseinate complex and restoring
the emulsifying properties of cheese proteins. The influence
of emulsifying salts on the final processed cheese structure
has been discussed by Kalab, Modler, Caric, and Milanovic
(1991) and Awad, Abdel-Hamid, El-Shabrawy, and Singh
(2002), among others; those salts also aid in controlling
pH and stability of disperse proteins, thus contributing to
the development of an appropriate microstructure.
Several reports are available pertaining to manufacture
of processed cheeses, using lower grade White, Cheddar,
Gruye`re, Gouda, Kashkaval, Emmental and Feta cheeses
(and other ingredients) as feedstock. For instance, Tamime,
Kalab, Davies, and Younis (1990) studied the processing of
Cheddar cheese with skim milk powder, and its effect upon
the microstructure of the final product. Kalab et al. (1991)
found, in turn, that White cheese is suitable to manufacture
processed cheese, as the original matrix is harder even at
high levels of cheese incorporation. Finally, Piska and
Stetina (2004) focused on cheese ripening and rate of
cooling of the blend.
Microbiological factors
In microstructural studies pertaining to Serra da Estrela
cheese, Parker, Gunning, Macedo, Malcata, and Brockle-
hurst (1998) found bacteria embedded in the protein matrix,
and especially lining the curd junctions; they also came
across with protein that appears as strands and free fat in
the smear e likely as a result of proteolytic and lipolytic
activities. Furthermore, cells were either directly in contact
with the fat globule membrane (as happened with most bac-
teria) or located at the caseinefat interface. On the other
hand, Lopez, Maillard, Briard-Bion, Camier, and Hannon
(2006) found bacteria organized as colonies in the matrix
of Emmental cheese e such colonies were preferentially
localized at the fat/protein interface, and both fat and bac-
teria were entrapped in the casein network; similar results
were previously obtained during characterization of Mozza-
rella cheese (Oberg, McManus, & McMahon, 1993).
Lactic acid bacteria have been claimed to contribute to
cheese microstructure, viz. Lactococcus lactis e via their
role in proteolysis (Centeno, Tomillo, Fernandez-Garca, Innovations
Examples
Gaya, & Nunez, 2002), and Lactobacillus casei e via its ca-
pacity to improve cheese body and texture (Merrill et al.,
Existing product
1996). Strains of Lactobacillus bulgaricus and Streptococ-
cus thermophilus have also been currently used in cheese
manufacture to increase moisture content and improve
Herbs/ham/etc. Incorporation of new ingredients
melting properties (Low et al., 1998; Peterson, Dave,
McMahon, Oberg, & Broadbent, 2000).
The hyphae of Penicillium camemberti, Penicillium Reduction/avoidance of harmful-like
Light product
ingredients
roqueforti, Penicillium glaucum or Mucor rasmussen pene-
trate the protein matrix, but their sporangia develop only
Bio product Incorporation of pre/probiotic ingredients
on the surface (Kalab, 1993); e.g. in Camembert cheese,
P. camemberti spores germinate on the cheese surface and
penetrate the curd, with a thick white mat of sporangia grow- Vitamins/Ca2+/ Inclusion of component(s) with nutraceutical
Peptides/etc. value
ing on the cheese surface (Rousseau, 1984). Brooker (1987)
found that deamination of amino acids by moulds on the Improvement/development of novel
Flavouring agents
surface increases pH, and consequently leads to precipita- flavours
tion of calcium phosphate from the aqueous phase e which,
in turn, produces a gradient between the surface and the bulk
of the cheese; this promotes a sandy mouthfeel, which is
New product
directly related to microstructure. Furthermore, such an
increase in pH facilitates the action of plasmin, thus contrib-
Fig. 1. Schematic formulation path of a novel dairy product.
uting further to cheese softening (Everett & Auty, 2008).
On the other hand, yeasts in cheese have been claimed
affects, in turn, development of microbial populations e
(Jakobsen & Narvhus, 1996) to assist in growth of P. roque-
which are implicated in flavour development or may even
forti by producing gas e which leads to curd openness; it thus
be vectors of intoxications. If microstructure issues are not
affects microstructure, besides supporting said growth e via
properly addressed, then technological improvement of
secretion of suitable nutrients thereinto.
foods will be commercially hampered by inadequate sensory
As a vector to protect bacteria from the putative
and safety features. Furthermore, incorporation in cheese of
unfriendly conditions found in cheese environments, immo-
bacteria that have proven in vitro to be beneficial to health
bilization of cells e via e.g. microencapsulation, appears to
requires adequate microenvironments e which are strongly
be a promising technique. According to O zer, Kirmaci,
dependent on the mutual effects of the proteinaceous net-
xenel, Atamer, and Hayaloglucth (2009), microencapsula-
S
work and the milk fat globule distribution.
tion does not adversely affect appearance or texture of
A new era in dairy food processing appears to be about to
experimental cheeses with immobilized bacteria, yet it
begin; the chief concern of industry e which is obviously
impacts significantly upon aroma and flavour attributes.
driven by consumer preference, will hereafter be coupled
with fundamental knowledge made available via sophisti-
Conclusions cated analytical apparata and imaging techniques; this will
At present, several food trends encompass creating desir- eventually lead to rational design of novel cheeses and ratio-
able, distinctive or novel textures, framed by attractive nal improvement of existing ones. Further complementary
appearance and appealing odour and taste. In particular, studies on microstructure subjects are thus fully warranted.
innovation in the dairy field relies nowadays mainly on: i) in-
corporation of nonconventional ingredients; ii) reduction in
content, or avoidance of harmful-like ingredients; iii) Acknowledgments
incorporation of pre- or/and probiotic vectors; iv) inclusion Financial support for author C. I. P. e provided via
of components with nutraceutical value; and v) enhancement a PhD fellowship (ref. SFRH/BD/18258/2004), issued by
of existing, or development of alternative flavours (Fig. 1). program POCI 2010, supervised by author F. X. M. and
Consumers are indeed calling for improvements in the sec- administered by Fundac~ao para a Ciencia e a Tecnologia
ondary (or organoleptic, including texture perception) role (Portugal), is hereby gratefully acknowledged.
of foods, but are simultaneously becoming more and more
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